Does rapid utilization of elevated nutrient availability allow eucalypts to dominate in the tropical savannas of Australia?

Northern Australia's savannas are among the most fire‐prone biomes on Earth and are dominated by eucalypts (Eucalyptus and Corymbia spp.). It is not clear what processes allow this group to dominate under such extreme fire frequencies and whether a superior ability to compete for nutrients and water might play a role. There is evidence that eucalypts are adapted to frequent fires; juvenile eucalypts escape the fire trap by growing rapidly in height between fires. However, non‐eucalypts are less able to escape the fire trap and tend to have stand structures strongly skewed toward suppressed juveniles. The mechanisms that drive these contrasting fire responses are not well understood. Here, we describe the results of a controlled glasshouse seedling experiment that evaluated the relative importance of nutrient and water availability in determining height growth and biomass growth of two eucalypt and one noneucalypt tree species, common in northern Australian savannas. We demonstrate that growth of eucalypt seedlings is particularly responsive to nutrient addition. Eucalypt seedlings are able to rapidly utilize soil nutrients and accumulate biomass at a much greater rate than noneucalypt seedlings. We suggest that a seasonal spike in nutrient availability creates a nutrient‐rich microsite that allows eucalypt seedlings to rapidly gain height and biomass, increasing their likelihood of establishing successfully and reaching a fire‐resistant size. Our results extend our understanding of how eucalypts dominate northern Australian savannas under extremely high fire frequencies.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Plants,fungi and bettongs: A fire-dependent co-evolutionary relationship

Digging activity by Bettongia gaimardi increased in small areas burnt by a low intensity fire at a site in northern Tasmania. A gap of several days occurred after the fire before digging began. Sporocarps of only one genus of mycorrhizal fungi (Mesophellia) were eaten by bettongs after the fire, in contrast to high numbers of species eaten in surrounding unburnt areas. It is hypothesized that the fire selectively stimulated the production of sporocarps by Mesophellia (a genus associated with eucalypts) rather than by simply making them more accessible or detectable. Stimulation of production of sporocarps by fire may ensure high infection rates and facilitate the uptake by eucalypts of nutrients released by the fire. These observations have important management implications.     

Bark thickness determines fire resistance of selected tree species from fire-prone tropical savanna in north Australia

We investigated the fire resistance conferred by bark of seven common tree species in north Australian tropical savannas. We estimated bark thermal conductance and examined the relative importance of bark thickness, density and moisture content for protecting the cambium from lethal fire temperatures. Eucalypt and non-eucalypt species were contrasted, including the fire-sensitive conifer Callitris intratropica. Cambial temperature responses to bark surface heating were measured using a modified wick-fire technique, which simulated a heat pulse comparable to surface fires of moderate intensity. Bark thickness was a better predictor of resistance to cambial injury from fires than either bark moisture or density, accounting for 68% of the deviance in maximum temperature of the cambium. The duration of heating required to kill the cambium of a tree (τ_c) was directly proportional to bark thickness squared. Although species did not differ significantly in their bark thermal conductance (k), the thinner barked eucalypts nevertheless achieved similar or only slightly lower levels of fire resistance than much thicker barked non-eucalypts. Bark thickness alone cannot account for the latter and we suggest that lower bark moisture content among the eucalypts also contributes to their apparent fire resistance. Unique eucalypt meristem anatomy and epicormic structures, combined with their bark traits, probably facilitate resprouting after fire and ensure the dominance of eucalypts in fire-prone savannas. This study emphasises the need to take into account both the thermal properties of bark and the mechanism of bud protection in characterising the resprouting ability of savanna trees.     

Groundwater rise,soil salinization and the decline of Casuarina in southeastern Australia during the late Quaternary

Abstract The decline in interglacial importance of Casuarina over the late Quaternary across southeastern Australia is documented. Three previously proposed causes for the decline (change in fire regime, change to a wetter climate and competitive exclusion by eucalypts) are shown to be inadequate for explaining the majority of cases. Re-examination of the evidence shows the decline at most sites occurred synchronously with a rise in groundwater or soil salinization, or both. From a review of biological literature, it is established that Casuarina stricta, the main species considered to have been affected by the decline, is likely to be disadvantaged by high water tables and saline soils. A link is demonstrated between groundwater salinity and the nodulation status of Casuarina in Victoria. It is concluded that the late Quaternary Casuarina decline was caused by a combination of rising groundwater levels and soil salinization. Soil salinization and groundwater level must therefore be considered as major factors determining vegetation patterns in southeastern Australia through the Quaternary up to the present day.     

The impact of experimental fire regimes on seed production in two tropical eucalypt species in northern Australia

Abstract This study investigated the effect of three experimental fire regimes on the fecundity, ovule development and seedfall of two common wet-dry tropical savanna eucalypts, Eucalyptus minima and Eucalyptus tetrodonta, in northern Australia. Both species flower early in the dry season and ovule development occurs during the dry season. This coincides with a period of frequent fires. The three fire regimes considered were applied for four years between 1990 and 1994. These regimes were (i) Unburnt, (ii) Early, fires lit early in the dry season, and (iii) Late, fires lit late in the dry season. The treatments were applied to nine catchments (15–20 km²) with each fire regime replicated three times. Fire intensity typically increases as the dry season proceeds. Therefore, early dry season fires generally differ from late dry season fires in both their intensity and their timing in relation to the reproductive phenology of the eucalypts. Late dry season burning significantly reduced the fecundity of both species, whereas Early burning had no significant effect. Ovule success was significantly reduced by the Early burning for both species. The Late burning significantly reduced ovule success in E. tetrodonta, but not in E. miniata. The results suggest that fire intensity and fire timing may both be important determinants of seed supply. Fire intensity may be a determinant of fecundity, whereas fire timing in relation to the reproduction phenology may have a significant impact on ovule survival. Both fire regimes resulted in a substantial reduction in seed supply compared with the Unburnt treatment. This may have a significant impact on seedling regeneration of these tropical savanna eucalypts.     

Interactions between fire,mycophagous mammals,and dispersal of ectromycorrhizal fungi in Eucalyptus forests

Several species of marsupials in Eucalyptus forests in Australia feed predominantly on the sporocarps of hypogeous fungi. This feeding is apparently beneficial to the fungi as it results in dispersal of spores. As these fungi are in almost all cases ectomycorrhiza-forming species, mycophagy by mammals may play an important role in the maintenance of mycorrhizal symbiosis in Eucalyptus forests. Fire is frequent and a dominant ecological factor in these forests, and this study tested the hypothesis that fire triggers both increased sporocarp production by some hypogeous ectomycorrhizal fungi associated with eucalypts, and increased mycophagy by mammals. Three experimental burns were set in E. tenuiramus forest in southeastern Tasmania. Digging activity (which reflects feeding on hypogeous fungi) by a mycophagous marsupial, the Tasmanian bettong Bettongia gaimardi, increased up to ten-fold after fire, with a peak about 1 month post-fire. This was associated with a similar pattern of increase in sporocarp production, which was due to species in the family Mesophelliaceae (especially Castoreum tasmanicum and Mesophellia spp.). This family appears to have radiated in association with eucalypts and has an exclusively Australasian distribution, unlike many of the other ectomycorrhizal fungi collected in this study which are cosmopolitan and have broad host ranges. No B. gaimardi were killed by fire, and there was no increase in mortality following fire. Population density increased after fire as a result of immigration of adult males. However, body condition and fecundity of individual B. gaimardi were maintained at pre-fire levels. This suggests that the availability of energy to B. gaimardi increased as a result of fire, and the fact that the contribution of fungus to the diet of B. gaimardi was high on burnt relative to control sites suggests further that this increase in energy availability was provided by hypogeous fungi. Effects of fire on hypogeous fungi and B. gaimardi were short-lived; all measured variables returned to control values about 4 months after fire. The capacity of B. gaimardi to survive fire and to harvest the increased sporocarp production triggered by fire provides a mechanism for the rapid dispersal of spores after fire. This should result in the establishment of ectomycorrhizae very early in post-fire succession. Because only some species of ectomycorrhizal fungi fruited in response to burning, fire probably has a strong influence on community structure among ectomycorrhizal fungi.     

Time‐since‐fire and climate interact to affect the structural recovery of an Australian semi‐arid plant community

How does time‐since‐fire influence the structural recovery of semi‐arid, eucalypt‐dominated Murray‐Mallee shrublands after fire, and is recovery affected by spatial variation in climate? We assessed the structure and dynamics of a hummock grass, Triodia scariosa N.T. Burb, and mallee eucalypts – two key structural components of mallee shrublands – using a >100 year time‐since‐fire chronosequence. The relative influence of climatic variables, both individually and combined with time‐since‐fire, was modelled to account for spatial variation in the recovery of vegetation structural components. Time‐since‐fire was the primary determinant of the structural recovery of T. scariosa and eucalypts. However, climate, notably mean annual rainfall and rainfall variability, also influenced the recovery of the eucalypt overstorey, T. scariosa cover and mean hummock height. We observed that (i) the mean number of live eucalypt stems per individual decreased while mean individual basal area increased, (ii) cover of T. scariosa peaked at ~30 years post‐fire and gradually decreased thereafter, and (iii) the ‘hummock’ form of T. scariosa occurred throughout the chronosequence, whereas the ‘ring’ form tended not to occur until ~30 years post‐fire. Time‐since‐fire was the key determinant of the structural recovery of eucalypt‐dominated mallee shrublands, but there is geographical variation in recovery related to rainfall and its variability. Fire regimes are likely to have different effects across the geographic range of mallee shrublands.     

Operculum development in theEudesmieae B eucalypts andEucalyptus caesia (Myrtaceae)

In theEudesmieae B eucalypts and inEucalyptus caesia, the perianth of the mature flower consists of a single, anatomically continuous, opercular structure that is crowned by the tips of the original free and separate calycine and corolline whorls. Ontogenetic and comparative evidence supports the hypothesis that this operculum is mostly corolline in composition, and that the calycine parts have been elevated distally onto the dorsal surface. In theEudesmieae B eucalypts this condition appears to be due to precocious initiation of the corolline primordia, followed by expansion and continuity of their growth centres which incorporates the areas at or below the base of the still differentiating calycine whorl. InEucalyptus caesia the corolline primordia are not precocious, but a similar situation is effected by a seemingly retarded increase in receptacle diameter relative to lateral expansion of the corolline growth centre(s). In these two examples the same final perianth form apparently derives from two different sets of growth processes which, nonetheless, result in the same allometric relationships within the developing flower. Although identical at maturity, the operculum form in these two taxa is best described in terms of convergence, rather than homology.     

Is decline in high altitude eucalypt forests related to rainforest understorey development and altered soil bacteria following the long absence of fire?

Abstract The objective of this study was to identify attributes of the understorey vegetation, soil root biomass, soil chemistry and microbial community that may be associated with tree decline in high altitude eucalypt forests in Tasmania. The sites studied were in healthy eucalypt forest, forest in decline and forest containing dead eucalypts dominated by rainforest, in north‐east (Eucalyptus delegatensis forest) and in north‐west (Eucalyptus coccifera forest) Tasmania. In both regions bare ground, rock and shrubby species were associated with healthy sites whereas decline sites were associated with moss and a tall understorey with a high percentage cover of rainforest species. Healthy sites had low root biomass in the top 10 cm of the soil profile relative to decline and rainforest sites. Seedlings of high altitude species were grown in rainforest soil (0.314% N and 0.060% P) and healthy eucalypt soil (0.253% N and 0.018% P). The four eucalypt species studied had similar root to shoot ratio in the two soils, but the rainforest species, Nothofagus cunninghamii and Leptospermum lanigerum, had higher root to shoot ratio in the healthy eucalypt than in the rainforest soil. We produced three soil filtrates: (i) fungi and bacteria present; (ii) bacteria only present and; and (iii) sterile, from healthy, decline and rainforest sites in north‐east and in north‐west Tasmania and used linseed as a germination bioassay. Filtrates from the north‐east decline and rainforest sites induced a significantly greater dysplastic germination response than healthy sites in (i) and (ii) filtrates, but this was not found in filtrates from sites in the north‐west. We conclude that while the development of a rainforest understorey and elevated soil root biomass in the long absence of fire is generally associated with high altitude eucalypt decline, altered bacterial and/or chemical attributes of soil are not always associated with high altitude eucalypt decline.     

Influence of fire severity on the regeneration,recruitment and distribution of eucalypts in the Cotter River Catchment,Australian Capital Territory

Abstract Plant responses to fire are variable between and within species and are influenced by numerous factors including fire severity. This study investigated the effects of fire severity on the regeneration and recruitment of forest eucalypts in the Cotter River Catchment, Australian Capital Territory (ACT). This study also examined the potential for the obligate seeder Eucalyptus delegatensis R.T. Baker (Myrtaceae) to expand into adjacent stands dominated by the facultative resprouter Eucalyptus fastigata H. Deane & Maiden (Myrtaceae) by seed shed and seedling establishment beyond the pre‐fire boundary. Sites were located in areas of either higher or lower fire severity, and transects were placed across the boundary of stands of E. delegatensis and E. fastigata. Species distributions, tree survival and seedling densities and heights were recorded, and the location of each boundary was determined as the region of maximum change in species composition along the transects. Eucalyptus delegatensis was the only eucalypt killed by higher severity fire. However, E. delegatensis seedling density was greater at higher severity sites than lower severity sites. Eucalyptus fastigata seedling density was low across all sites, with other eucalypts producing few, if any, seedlings. There was no evidence that E. delegatensis had increased its range into downslope stands dominated by E. fastigata. Patterns of vegetative recovery and seedling recruitment may be related to a number of factors, including differences in allocation patterns between seeders and sprouters, and the effects of overstory and understory competition. It is unclear what processes impede E. delegatensis seedling establishment beyond the stand boundary, but may involve an inability of E. delegatensis to shed seed sufficiently far downslope; unsuitable conditions for germination beyond the boundary; or, competition from a retained or resprouting overstory, despite the potential for increased dispersal distance soon after fire.     

Premature Decline of <Emphasis Type="Italic">Eucalyptus</Emphasis> and Altered Ecosystem Processes in the Absence of Fire in Some Australian Forests

We propose a model of ‘premature tree decline’ whereby an absence of fire hastens the mortality of overstorey eucalypts in some forests. This model is relevant to some temperate Australian forests in which fire regimes have shifted from relatively frequent before European settlement to infrequent following settlement. The increased development of midstorey vegetation and litter accumulation has occurred since European settlement in some specific examples of Australian forests and woodlands. Our model proposes that in the long absence of fire: 1. midstorey vegetation reduces the availability of soil water for eucalypts and; 2. Eucalypts have less access to P and/or cations as these elements become locked up in soil, litter and midstorey biomass. We highlight important knowledge gaps and argue that research into ecological burning, for eucalypt health and other values such as biodiversity, is urgently required.     

A conceptual model of vegetation dynamics for the unique obligate‐seeder eucalypt woodlands of south‐western Australia

Models of vegetation dynamics framed as testable hypotheses provide powerful tools for predicting vegetation change in response to contemporary disturbances or climate change. Synthesizing existing information and applying new data, we develop a conceptual model of vegetation states and transitions for the previously overlooked woodlands dominated by obligate‐seeder eucalypts of dry to semi‐arid south‐western Australia. These comprise the largest extant temperate woodland globally, are uniquely dominated by a high diversity of obligate‐seeder eucalypts (55 taxa), but are under threat from wildfire. Our conceptual model incorporates four critical ecological processes that also distinguish obligate‐seeder woodlands from temperate woodlands dominated by resprouting eucalypts: (i) a lack of well‐protected epicormic buds results in major disturbances (prominently fire) being stand‐replacing. The pre‐disturbance tree cohort is killed, followed by dense post‐disturbance recruitment from seed shed from a serotinous seed bank; (ii) competition between saplings leads to self‐thinning over a multi‐century timeframe, with surviving individuals having great longevity (regularly >400 years); (iii) multiple processes limit recruitment in the absence of stand‐replacement disturbances, leading to frequent single‐cohort stands. However, unlike the few other obligate‐seeder eucalypt communities, trickle recruitment in very long‐unburnt stands can facilitate indefinite community persistence in the absence of stand‐replacement disturbances; and (iv) discontinuous fuels, relatively low understorey flammability (low grass and often high chenopod cover) and topographic barriers to fire (salt lakes) allow mature woodlands to persist for centuries without burning. Notably though, evidence suggests that flammability peaks at intermediate times since fire, establishing a ‘flammability bottleneck’ (or landscape fire trap) through which regenerating woodlands must pass. Our model provides a framework to support management to conserve obligate‐seeder eucalypt woodlands. Research into reasons for exceptional tree heights relative to ecosystem productivity, the evolution of diverse and dominant obligate‐seeder eucalypts, the paucity of grass, and the recent spatial distribution of fires, will further inform conservation management.     

Demographic changes and the development of competition in a native Australian eucalypt forest invaded by exotic pines

Data from the second half of a 14-year study of a eucalypt dry sclerophyll forest invaded by Pinus radiata D. Don shows further progress in the development of a mixed eucalypt-pine stand. Earlier (Chilvers and Burdon 1983) it was clear that pines invading from an adjacent commercial plantation were starting to contribute their own progeny to the mixed stand. Initially the recruitment of established pines (> 1 m high) from these seedlings was very slow. However, since 1984, increasing numbers of seedlings have survived beyond their first year, suggesting the possibility of another wave of successful establishment in the near future. The established fraction of the pine population continued to grow rapidly and incurred no mortality. Many of these plants have now emerged through the top of the eucalypt canopy. In contrast, both juvenile and adult eucalypt numbers continued to decline. Despite these demographic and visual changes, comparative analysis of the growth of established pines over two consecutive 7-year periods (1974–1981; 1981–1988) showed that competition between neighbouring plants, especially pines, is beginning to affect individual growth rates. Growth rates of pines having other pines as nearest neighbours declined 40% between 1974–1981 and 1981–1988, while that of pines with eucalypts as nearest neighbours fell by 28%. In pine-pine nearest neighbour (NN) pairs, growth rates were significantly affected by the size (diameter) of the nearest neighbour. In pine-eucalypt NN pairs growth rates of the pines were not affected by the size of the eucalypt, but these were significantly negatively correlated with the inverse of the distance to the eucalypt. More broad-based assessment of 43 pines showed a significant relationship between their growth rates and the summed competitive effect of other pines within a 5-m radius. No such relationship was found between growth rate and the summed competitive effect of eucalypts in the 5-m zone.     

Impact of fire on leaf nutrients,arthropod fauna and herbivory of native and exotic eucalypts in Kings Park,Perth, Western Australia

The vegetation of Kings Park, near the centre of Perth, Western Australia, once had an overstorey of Eucalyptus marginata (jarrah) or Eucalyptus gomphocephala (tuart), and many trees still remain in the bushland parts of the Park. Avenues and roadsides have been planted with eastern Australian species, including Eucalyptus cladocalyx (sugar gum) and Eucalyptus botryoides (southern mahogany), both of which have become invasive. The present study examined the effect of a recent burn on the level of herbivory on these native and exotic eucalypts. Leaf damage, shoot extension and number of new leaves were measured on tagged shoots of saplings of each tree species in unburnt and burnt areas over an 8‐month period. Leaf macronutrient levels were quantified and the number of arthropods on saplings was measured at the end of the recording period by chemical knockdown. Leaf macronutrients were mostly higher in all four species in the burnt area, and this was associated with generally higher numbers of canopy arthropods and greater levels of leaf damage. It is suggested that the pulse of soil nutrients after the fire resulted in more nutrient‐rich foliage, which in turn was more palatable to arthropods. The resulting high levels of herbivory possibly led to reduced shoot extension of E. gomphocephala, E. botryoides and, to a lesser extent, E. cladocalyx. This acts as a negative feedback mechanism that lessens the tendency for lush, post‐fire regrowth to outcompete other species of plants. There was no consistent difference in the levels of the various types of leaf damage or of arthropods on the native and the exotic eucalypts, suggesting that freedom from herbivory is not contributing to the invasiveness of the two exotic species.     

Responses of forest eucalypts to moderate and high intensity fire in the Tingle Mosaic,south‐western Australia: comparisons between locally endemic and regionally distributed species

Responses of three locally endemic (Eucalyptus brevistylis, Eucalyptus jacksonii and Eucalyptus guilfoylei) and three co‐occurring regional eucalypts (Eucalyptus marginata, Eucalyptus diversicolor and Corymbia calophylla) to moderate‐ and high‐intensity fires were examined in granitic terrain of the Tingle Mosaic, south‐western Australia. Significant associations between diameter distributions and community type (CT) for each species (P < 0.001) suggest that fire response will also vary according to the habitat/fire interaction. None of the species were fire sensitive, although responses differed both within and between species, and with CT. All species examined predominately consisted of several cohorts of regeneration within a forest stand. Each species had thick bark and re‐sprouted from crown epicormics following 100% scorch of the mature tree. The quantity and type of regeneration in relation to gaps created by individual dead trees following fire differed between species; for example, E. guilfoylei regeneration was strongly associated with gaps, and C. calophylla with non‐gaps. However, regeneration of the two tall open‐forest species, E. jacksonii and E. diversicolor were not most associated with either gaps or non‐gaps. The very low levels of regeneration of E. brevistylis following fire and the high proportion of stems of E. jacksonii that were hollow butted (40% of stems > 1 m DBHOB) may be factors associated with narrow endemism of these species and may affect the vulnerability of these eucalypts to fire. The interaction of seed availability, intense fires and subsequent rainfall may be critical in the long term survival of these species. Eucalyptus guilfoylei, by contrast, appears well adapted to the increasing levels of disturbance likely in the region where these species occur. The vulnerability of a locally endemic species in a fire‐prone environment is likely to reflect differences to the prevailing adaptations of the dominant species rather than an inherent ability of the species to survive or respond. Management regimes must account for variations in species responses to fire in different CTs if the long‐term survival chances of local endemics are to be enhanced.     

Eucalyptus regnans (Myrtaceae): A fire-sensitive eucalypt with a resprouter epicormic structure

Determining the location of buds and bud-forming meristems and hence the level of protection from heat is essential to understanding plant response to fire. Most eucalypts resprout readily from the stem (epicormic resprouting) and the base after felling or high intensity fire. In contrast, Eucalyptus regnans is one of the few eastern Australian fire-sensitive, obligate seeder eucalypts. Some authors have suggested that the relatively weak epicormic resprouting is due to a lack of bud-forming structures. Epicormic strands from the bark and outer xylem of three very large trees and two saplings were examined anatomically. Epicormic bud-forming structures were found in all samples examined. The bud-forming capacity consisted of narrow, radially elongated strips of cells of meristematic appearance. These strips were continuous from the outermost secondary xylem through to the outer bark. Bark was relatively thick at the base of the large trees, but remarkably thin above this basal skirt. Eucalyptus regnans was found to possess the apparently fire-adapted epicormic strands previously described in other eucalypts, thus showing its fire-adapted lineage. However, this fire-sensitive species apparently directs much of its resources to rapid height-growth rates in younger trees, rather than to vegetative fire survival.     

Contributions of insects and droughts to growth decline of trembling aspen mixed boreal forest of western Canada

Insects, diseases, fire and drought and other disturbances associated with global climate change contribute to forest decline and mortality in many parts of the world. Forest decline and mortality related to drought or insect outbreaks have been observed in North American aspen forests. However, little research has been done to partition and estimate their relative contributions to growth declines. In this study, we combined tree‐ring width and basal area increment series from 40 trembling aspen (Populus tremuloides Michx.) sites along a latitudinal gradient (from 52° to 58°N) in western Canada and attempted to investigate the effect of drought and insect outbreaks on growth decline, and simultaneously partition and quantify their relative contributions. Results indicated that the influence of drought on forest decline was stronger than insect outbreaks, although both had significant effects. Furthermore, the influence of drought and insect outbreaks showed spatiotemporal variability. In addition, our data suggest that insect outbreaks could be triggered by warmer early spring temperature instead of drought, implicating that potentially increased insect outbreaks are expected with continued warming springs, which may further exacerbate growth decline and death in North America aspen mixed forests.     

Are the eucalypt and non-eucalypt components of Australian tropical savannas independent?

Eucalypts (Eucalyptus and Corymbia spp.) dominate (>60%) the tree biomass of Australia's tropical savannas but account for only a fraction (28%) of the tree diversity. Because of their considerable biomass and adaptation to environmental stressors, such as fire, the eucalypts may drive tree dynamics in these savannas, possibly to the exclusion of non-eucalypts. We evaluated whether the eucalypt and non-eucalypt components in tropical savannas are dependent so that changes in one component are matched by opposite trends in the other. Using tree inventory data from 127 savanna sites across the rainfall and fire frequency gradients, we found that eucalypt and non-eucalypt basal area and species richness had a negative relationship. This relationship was maintained across the rainfall gradient, with rainfall having a positive effect on the basal area and species richness of both components, but with a greater effect in non-eucalypts. Fire frequency negatively affected basal area, but not species richness, although basal area and species richness of eucalypts and non-eucalypts did not differ in their response to fire. Rainfall appears to set the upper bounds to woody biomass in these mesic savannas, while fire maintains woody biomass below carrying capacity and facilitates coexistence of the components. The magnitude of the component responses, particularly for non-eucalypts, is determined by rainfall, but their dependence is likely due to their differential response to both rainfall and fire, but not to competition for resources. Thus, while eucalypts dominate biomass overall, at high rainfall sites non-eucalypt basal area and diversity are highest, especially where fire frequency is low.     

On the ecological role of insects in Australian eucalypt forests

If heterotroph grazers act as rate regulators of autotroph processes, the general intensity of grazing probably reflects the necessity for more or less intense rate-regulation. The high rates of insect grazing in eucalypt forests imply that considerably more regulation of eucalypts is necessary than in other forests. After examining the biological adaptations of eucalypts and reviewing nutrient cycling in eucalypt forests, the hypothesis is proposed that the impoverished soils and erratic Australian climate have led to the development of very efficient autotrophic exploitation of the resource base, with associated sensitive and rapid growth responses to favourable conditions and a correspondingly sensitive and intense level of insect grazing. The rate-regulatory activity of the insects ensures that the resource base is not depleted, and it maintains a proportion of the available nutrients in rapid circulation. In addition to the observed pattern of intense insect grazing in eucalypt forests, it is suggested that a two-pathway nutrient cycling system has been developed, with pathways more clearly defined than the anastomosing nutrient-cycling systems in forests elsewhere. A slow nutrient cycle is maintained by the trees while a smaller pool of nutrients is rapidly recycled in the litter and understorey. Insects skim off nutrients from the slow cycle and pass them into the faster cycling pool. This idea is important in relation to the frequency of fire in eucalypt forests and the large inputs into the system of nitrogen from the leguminous shrubs which regenerate after fire.